IntroductionProteins of the extracellular matrix in animal cells or the cell wall in plant cells have only evolved after the divergence of plants and animals, about 1 billion years ago during the Precambrian. Although the cell walls of seed plants are mainly composed of polysaccharides such as cellulose, hemicelluloses and pectins, they also contain variable amounts of structural proteins. These proteins have traditionally been classified as glycine-rich proteins (GRPs), proline-rich proteins (PRPs) and hydroxyproline-rich proteins (HRGPs). With the exception of GRPs, they are glycosylated and contain hydroxyproline (Carpita and Gibeaut, 1993;Cassab, 1998;Zablackis et al., 1995). Dividing and elongating plant cells contain very little cross-linked structural cell-wall proteins. As a rule, cell-typeand tissue-specific cross-linking of these proteins into the cell wall occurs towards the end of elongation growth. Little is known about the roles of these proteins. It is thought that they strengthen the cell walls in normal plant development (Fry, 1988;Carpita and Gibeaut, 1993) and are involved in incompatible plant-pathogen interactions (Lamb and Dixon, 1997;Delledonne et al., 1998).Xylem is the principal water-conducting tissue of vascular plants. It contains water pipes formed by the walls of dead cells. To allow the efficient transport of water and solutes, the end walls of the cylindrical cells are perforated (in vessel members) or the ends are elongated and make contacts side by side (in tracheids) by numerous pit pairs. Vessel members are absent from most gymnosperms but are typically observed in the wood of angiosperms. However, the xylem of elongating organs of angiosperms often contains both cell types. For convenience, they are collectively called 'tracheary elements ' (Esau, 1977). The main steps of the differentiation of tracheary elements are cell elongation, strengthening of the wall by deposition of secondary wall layers, lignification and programmed cell death initiated by the collapse of the vacuole and followed by the degradation of cellular constituents by vacuolar hydrolases (Groover et al., 1997;Groover and Jones, 1999;Fukuda, 2000). Finally, the hydrolysis products are probably taken up by the living xylem parenchyma surrounding the tracheary elements.Protoxylem is the water-conducting tissue of young and elongating plant organs, whereas metaxylem starts to differentiate only towards the end of elongation growth. The challenge for the growing plant is to maintain a functional water-conducting system made up of the cell walls of dead protoxylem elements (PX) embedded in rapidly elongating xylem parenchyma cells Supplemental data available online